Field evidence for density-dependent facilitation amongst Ixodes ricinus ticks feeding on sheep.

Engorged Ixodes ricinus nymphs collected from sheep resident in an upland UK field site were significantly lighter than nymphs that engorged on previously tick-naïve sheep, indicating that site-resident sheep continually exposed to ticks acquired anti-tick resistance. The weights of engorged nymphs that fed on naturally tick-resistant sheep increased significantly, however, when increasingly high numbers of adult female ticks fed on the sheep during seasonal peaks of tick activity. This relationship was unaffected by variations in nymph weight amongst individual sheep, between seasons and years, and potential effects of sheep infection with Ehrlichia phagocytophila; this suggests that high adult tick infestations may directly inhibit the expression of acquired anti-tick resistance by sheep. The length, width and weight of adult ticks and the scutum length of adult females were linearly related to their weight as an engorged nymph. The mean scutum length of adult female ticks feeding on sheep in the field site was greater than that of adult females obtained from engorged nymphs collected from sheep of the same site. This suggests that larger ticks have a survival advantage and that I. ricinus ticks exhibit density-dependent intraspecific facilitation at high infestation levels with potential consequences for the transmission of tick-borne diseases.